Key to the females of Afrotropical Anopheles mosquitoes (Diptera: Culicidae)

1Dichotomous keys for the morphological identification of groups of organisms have been used for over 300 years. These keys lead the reader through a series of couplets, each one giving two choices of characters, until a species name is reached. For the anopheline mosquitoes of the Afrotropical Region, which includes some of the most efficient transmitters of malaria parasites in the world, the first key for their identification was published in 1931, and the most recent printed version in 1987, this last being a pictorial key containing line graphics of characters mentioned in each couplet. In the 32 years that have passed since Gillies and Coetzee published their key, several new species have been discovered, described and named.

2The pictorial key to adult female anophelines in the Afrotropical Region was used as the template for the current key. New illustrations were produced and new couplets inserted where appropriate to accommodate new species described since 1987.

3The present key is a revision of that presented in Gillies and Coetzee, with the addition of newly described species and the exclusion of subspecies. Table 1 provides a lists of species, authorship and current classification, while Table 2 gives the number of species described per decade since 1900. The user is encouraged to refer to both the 1968 volume of Gillies and De Meillon and the 1987 supplement of Gillies and Coetzee for full species descriptions, biology and geographic distribution. More recent references include Sinka et al., and Kyalo et al..

4The following species are not included in the keys: Anopheles ethiopicus lacks hindlegs, Anopheles erythraeus and Anopheles dualaensis adults are unknown, and Anopheles eouzani lacks hindtarsomeres 4 and 5. The following new species are included: Anopheles okuensis (Section I), Anopheles hervyi, Anopheles millecampsi and Anopheles multicinctus (Section IV), Anopheles rageaui and Anopheles seretsei (Section VII), Anopheles kosiensis (Section IX), Anopheles gabonensis (Section X), and Anopheles carnevalei and Anopheles ovengensis (Section XI).

5A major addition to the key is the inclusion of Anopheles stephensi (Section IV), the Asian malaria vector with distribution from the Middle East to China. This species was first detected on the African continent in Djibouti in September 2012 and subsequently in February 2013 through to December 2017. It has also recently been found in Ethiopia in 2016. The species is similar to those belonging to the Anopheles gambiae complex—mosquitoes with speckled legs—but differs by having the wing with two pale spots in the 2nd main dark area of the costa and vein 1, thus being similar to Anopheles maculipalpis and Anopheles pretoriensis, from which it differs by not having hindtarsomeres 4 and 5 all pale.

6There are several groups of species where morphological identification is not possible using only the adult females, either because the adults look identical or because of overlap in morphological variation. Some of these species can be identified on immature characters, thus requiring eggs or larvae, while others require genetical methods, such as chromosomal inversions or molecular assays. Such groups include:the well-known Anopheles gambiae complex (An. gambiae, Anopheles coluzzii, Anopheles arabiensis, Anopheles quadriannulatus, Anopheles amharicus, Anopheles fontenillei, Anopheles bwambae, Anopheles melas, Anopheles merus);the Anopheles funestus group (Anopheles funestus, An. funestus-like, Anopheles parensis, Anopheles vaneedeni, Anopheles aruni, Anopheles confusus, Anopheles leesoni, Anopheles rivulorum, An. rivulorum-like, Anopheles brucei, Anopheles fuscivenosus);the Anopheles nili complex (An. nili, Anopheles somalicus);the Anopheles marshallii complex (An. marshallii, Anopheles letabensis, Anopheles hughi, Anopheles kosiensis) and its allies Anopheles hargreavesi, Anopheles gibbinsi and Anopheles mousinhoi;Anopheles squamosus/cydippis, the former of which is known to consist of at least five chromosomal forms (Green and Hunt, unpublished data);Anopheles coustani/crypticus/namibiensis in southern Africa.

7the well-known Anopheles gambiae complex (An. gambiae, Anopheles coluzzii, Anopheles arabiensis, Anopheles quadriannulatus, Anopheles amharicus, Anopheles fontenillei, Anopheles bwambae, Anopheles melas, Anopheles merus);

8the Anopheles funestus group (Anopheles funestus, An. funestus-like, Anopheles parensis, Anopheles vaneedeni, Anopheles aruni, Anopheles confusus, Anopheles leesoni, Anopheles rivulorum, An. rivulorum-like, Anopheles brucei, Anopheles fuscivenosus);

9the Anopheles nili complex (An. nili, Anopheles somalicus);

10the Anopheles marshallii complex (An. marshallii, Anopheles letabensis, Anopheles hughi, Anopheles kosiensis) and its allies Anopheles hargreavesi, Anopheles gibbinsi and Anopheles mousinhoi;

11Anopheles squamosus/cydippis, the former of which is known to consist of at least five chromosomal forms (Green and Hunt, unpublished data);

12Anopheles coustani/crypticus/namibiensis in southern Africa.

13The definition of “complex”, as applied to the genus Anopheles, is a group of species that are virtually morphologically identical but are otherwise considered valid species. The use of the term “group” denotes species that are morphologically very similar at the adult stage but many can be distinguished at the immature stages.

14Except for those of medical importance, the above list is just a small sample of species groups about which we know very little biologically in terms of feeding/resting preferences or their role in malaria parasite transmission. Basic taxonomic research, aligned with molecular analyses, is still very much needed in the Culicidae.

15An updated key for the morphological identification of Afrotropical anopheline species is presented. This key should be used in conjunction with earlier works giving full species descriptions, biology, medical importance and distribution.

16Characters are presented in ‘couplets’ where two options are presented, giving two different outcomes, eventually ending at a species name. The illustration(s) for the first option of each couplet is on the left (or rarely, in the centre) and for the second option on the right. General terminology follows that of Harbach and Knight.

17Terminology of the wing venation has changed over the past 80 years in attempts to align Culicidae with the rest of the Diptera Family (Table 3). The terminology proposed in the recent Manual of Afrotropical Diptera has been challenged by culicid taxonomists (manuscript reviewers), specifically around the terms used for the posterior veins (veins 5 and 6 in Fig. 1). Since consensus on the terminology has not yet been reached, and given that the malaria vector control field workers in Africa have been using the Anopheles identification keys published 32 years ago that use the numbering shown in Fig. 1, this simplified system is used here. It avoids unnecessary repetition of the various terms in each couplet and reference can be made to Table 3 where recent terminology is required.

18Abdominal segments with laterally projecting tufts of scales on segments II–VII …Section IAbdominal segments not so…2 Hindtarsus with at least last 2 hindtarsomeres entirely pale …Section IIHindtarsus not so …3Hindtarsomere 5 mainly or entirely dark, hindtarsomere 4 white …Section IIIHindtarsus not so …4Legs speckled, sometimes sparsely …Section IVLegs not speckled …5Wing entirely dark or with pale spots confined to costa and vein 1 …Section VWing not so …6Wing without a pale spot on basal 0.5 of costa …Section VIWing with at least 1 pale spot on basal 0.5 of costa …7Maxillary palpus with apex dark …Section VIIMaxillary palpus with apex pale …8Maxillary palpus with 4 pale bands …Section VIIIMaxillary palpus with less than 4 pale bands …9Wing with pale interruption in 3rd main dark area (preapical dark spot) of vein 1, sometimes fused with preceding pale area …Section IX3rd main dark area without pale interruption …10Wing with 2 pale spots on upper branch of vein 5 …Section XWing with I pale spot on upper branch of vein 5 …Section XI

19Abdominal segments with laterally projecting tufts of scales on segments II–VII …Section I

20Abdominal segments not so…2

21Hindtarsus with at least last 2 hindtarsomeres entirely pale …Section II

22Hindtarsus not so …3

23Hindtarsomere 5 mainly or entirely dark, hindtarsomere 4 white …Section III

24Hindtarsus not so …4

25Legs speckled, sometimes sparsely …Section IV

26Legs not speckled …5

27Wing entirely dark or with pale spots confined to costa and vein 1 …Section V

28Wing not so …6

29Wing without a pale spot on basal 0.5 of costa …Section VI

30Wing with at least 1 pale spot on basal 0.5 of costa …7

31Maxillary palpus with apex dark …Section VII

32Maxillary palpus with apex pale …8

33Maxillary palpus with 4 pale bands …Section VIII

34Maxillary palpus with less than 4 pale bands …9

35Wing with pale interruption in 3rd main dark area (preapical dark spot) of vein 1, sometimes fused with preceding pale area …Section IX

363rd main dark area without pale interruption …10

37Wing with 2 pale spots on upper branch of vein 5 …Section X

38Wing with I pale spot on upper branch of vein 5 …Section XI

39Wing almost entirely dark, costa without pale spots …brumptiWing with abundant pale areas, costa with at least 4 pale spots …2Hindtarsomeres 1 to 5 entirely dark … argenteolobatus(southern Africa)murphyi (West Africa)Hindtarsomeres 1 to 4, at least, with apical pale bands …3Hindtarsomeres 1 and 2 with definite pale and dark rings in addition to apical pale bands …cinctusHindtarsomeres 1 and 2 with pale bands at apices only …4Hindtarsomeres 3 and 4 all white or narrowly dark basally, 5 all dark or at least basal 0.5 dark..5Hindtarsomeres not so …7Moderate-sized species; abdominal scale-tufts short and dark; 0.5 or more of hindtarsomere 1 pale …cristipalpisVery large species, abdominal segments II–VII with long lateral tufts of yellowish and dark scales; hindtarsomere 1 largely dark …6Pale fringe spot present opposite lower branch of vein 5 …okuensisNo pale fringe spot opposite lower branch of vein 5 …..implexusHindtarsomere 5 and about apical 0.5 of 4 pale …pharoensisHindtarsomere 5 all dark and 4 with much less than apical 0.5 pale …8Very small species (wing length 2.5–2.8 mm); wing with upper branch of vein 2 largely pale …swahilicusSmall to moderate-sized species (wing length 2.7–4.5 mm); wing with upper branch of vein 2 either entirely dark apart from apex or with a few scattered pale scales only …squamosus cydippis

40Wing almost entirely dark, costa without pale spots …brumpti

41Wing with abundant pale areas, costa with at least 4 pale spots …2

42Hindtarsomeres 1 to 5 entirely dark … argenteolobatus

43(southern Africa)

44murphyi

45(West Africa)

46Hindtarsomeres 1 to 4, at least, with apical pale bands …3

47Hindtarsomeres 1 and 2 with definite pale and dark rings in addition to apical pale bands …cinctus

48Hindtarsomeres 1 and 2 with pale bands at apices only …4

49Hindtarsomeres 3 and 4 all white or narrowly dark basally, 5 all dark or at least basal 0.5 dark..5

50Hindtarsomeres not so …7

51Moderate-sized species; abdominal scale-tufts short and dark; 0.5 or more of hindtarsomere 1 pale …cristipalpis

52Very large species, abdominal segments II–VII with long lateral tufts of yellowish and dark scales; hindtarsomere 1 largely dark …6

53Pale fringe spot present opposite lower branch of vein 5 …okuensis

54No pale fringe spot opposite lower branch of vein 5 …..implexus

55Hindtarsomere 5 and about apical 0.5 of 4 pale …pharoensis

56Hindtarsomere 5 all dark and 4 with much less than apical 0.5 pale …8

57Very small species (wing length 2.5–2.8 mm); wing with upper branch of vein 2 largely pale …swahilicus

58Small to moderate-sized species (wing length 2.7–4.5 mm); wing with upper branch of vein 2 either entirely dark apart from apex or with a few scattered pale scales only …squamosus

59cydippis

60Legs speckled …2Legs not speckled …8Hindtarsomeres 3 to 5 entirely pale …3Hindtarsomere 3 dark at base …5Maxillary palpus with 3 pale bands, usually with some speckling; vein 1 of wing with 2 pale spots in 2nd main dark area (median dark spot) … maculipalpisMaxillary palpus with 4 pale bands, unspeckled; vein 1 of wing with at most 1 pale spot in 2nd main dark area …4Midtarsomeres 2 to 4 entirely dark; vein 1 of wing dark at base, basal 0.5 of stem of vein 4 with small pale areas …maliensisMidtarsomeres 2 to 4 with pale apices; vein 1 of wing pale at base, basal 0.5 of stem of vein 4 entirely pale …deemingiHindtarsomere 1 broadly pale at apex; vein 1 of wing with 2 pale spots in 2nd main dark area …..pretoriensisHindtarsomere 1 narrowly pale or dark at apex; vein 1 of wing with 1 pale spot in 2nd main dark area ….6Foretarsomere 1 with 5–9 pale rings; stem of vein 4 of wing largely pale …machardyiForetarsomere 1 with 2–4 pale rings; stem of vein 4 of wing largely dark …7Fore- and midtarsomeres 2 and 3 pale at apex; wing with fringe spot opposite vein 6 …natalensisFore- and midtarsomeres 2 and 3 dark apically; no fringe spot opposite vein 6 …buxtoniMaxillary palpus very shaggy and unbanded or with 1–4 irregular narrow pale bands …9Maxillary palpus smooth with 3 pale bands, the 2 distal ones broad or rarely fused …14Maxillary palpus without pale bands; no pale spot at apex of hindtibia or base of hindtarsomere 1 …caliginosusMaxillary palpus with 1–4 pale bands; apex of hindtibia broadly or narrowly pale …10Hindtarsomere 3 entirely pale …11Hindtarsomere 3 dark at base …12Base of hindtarsomere 1 dark; pale fringe spot present opposite lower branch of wing vein 5 …paludisBase of hindtarsomere 1 broadly pale; no pale fringe spot opposite lower branch of wing vein 5 …coustani(in part)Hind tarsomere 1 entirely dark basally or at most with a very narrow band of pale scales not as broad as the width of the tarsomere …tenebrosus(in part)Hind tarsomere 1 broadly pale at base, pale area at least as long as broad …13Apex of hindtibia with a pale streak 3–5 times as long as broad; apical pale band on hindtarsomere 2 0.13–0.4 length of tarsomere …coustani(in part)crypticus (S. Africa only)Pale streak on hindtibia 1–3 times as long as broad; apical pale band on hindtarsomere 2 narrow, 0.07–0.13 length of segment ….ziemanninamibiensis 3rd main dark area on wing vein 1 without a pale interruption; foretarsomeres 1 to 3 usually without distinct apical pale bands..rufipes(in part)3rd main dark area on wing vein 1 with a pale interruption, or with a short extension of the subcostal pale spot into the dark area on vein 1; foretarsomeres 1 to 3 with apical pale bands …15Hindtarsomere 3 entirely pale.…hancockibrohieri (in part)Hindtarsomere 3 not so.…brohieri(in part) W. Africatheileri mainly E. & S. Africa

61Legs speckled …2

62Legs not speckled …8

63Hindtarsomeres 3 to 5 entirely pale …3

64Hindtarsomere 3 dark at base …5

65Maxillary palpus with 3 pale bands, usually with some speckling; vein 1 of wing with 2 pale spots in 2nd main dark area (median dark spot) … maculipalpis

66Maxillary palpus with 4 pale bands, unspeckled; vein 1 of wing with at most 1 pale spot in 2nd main dark area …4

67Midtarsomeres 2 to 4 entirely dark; vein 1 of wing dark at base, basal 0.5 of stem of vein 4 with small pale areas …maliensis

68Midtarsomeres 2 to 4 with pale apices; vein 1 of wing pale at base, basal 0.5 of stem of vein 4 entirely pale …deemingi

69Hindtarsomere 1 broadly pale at apex; vein 1 of wing with 2 pale spots in 2nd main dark area …..pretoriensis

70Hindtarsomere 1 narrowly pale or dark at apex; vein 1 of wing with 1 pale spot in 2nd main dark area ….6

71Foretarsomere 1 with 5–9 pale rings; stem of vein 4 of wing largely pale …machardyi

72Foretarsomere 1 with 2–4 pale rings; stem of vein 4 of wing largely dark …7

73Fore- and midtarsomeres 2 and 3 pale at apex; wing with fringe spot opposite vein 6 …natalensis

74Fore- and midtarsomeres 2 and 3 dark apically; no fringe spot opposite vein 6 …buxtoni

75Maxillary palpus very shaggy and unbanded or with 1–4 irregular narrow pale bands …9

76Maxillary palpus smooth with 3 pale bands, the 2 distal ones broad or rarely fused …14

77Maxillary palpus without pale bands; no pale spot at apex of hindtibia or base of hindtarsomere 1 …caliginosus

78Maxillary palpus with 1–4 pale bands; apex of hindtibia broadly or narrowly pale …10

79Hindtarsomere 3 entirely pale …11

80Hindtarsomere 3 dark at base …12

81Base of hindtarsomere 1 dark; pale fringe spot present opposite lower branch of wing vein 5 …paludis

82Base of hindtarsomere 1 broadly pale; no pale fringe spot opposite lower branch of wing vein 5 …coustani

83(in part)

84Hind tarsomere 1 entirely dark basally or at most with a very narrow band of pale scales not as broad as the width of the tarsomere …tenebrosus

85(in part)

86Hind tarsomere 1 broadly pale at base, pale area at least as long as broad …13

87Apex of hindtibia with a pale streak 3–5 times as long as broad; apical pale band on hindtarsomere 2 0.13–0.4 length of tarsomere …coustani

88(in part)

89crypticus

90(S. Africa only)

91Pale streak on hindtibia 1–3 times as long as broad; apical pale band on hindtarsomere 2 narrow, 0.07–0.13 length of segment ….ziemanni

92namibiensis

933rd main dark area on wing vein 1 without a pale interruption; foretarsomeres 1 to 3 usually without distinct apical pale bands..rufipes

94(in part)

953rd main dark area on wing vein 1 with a pale interruption, or with a short extension of the subcostal pale spot into the dark area on vein 1; foretarsomeres 1 to 3 with apical pale bands …15

96Hindtarsomere 3 entirely pale.…hancocki

97brohieri

98(in part)

99Hindtarsomere 3 not so.…brohieri

100(in part) W. Africa

101theileri

102mainly E. & S. Africa

103Femora and tibiae speckled …kingiFemora and tibiae with at most apical bands only …2Maxillary palpus shaggy; costa and vein 1 of wing without usual main dark areas …symesiMaxillary palpus smooth; 2nd main dark area of wing vein 1 well defined and with 2 pale interruptions …rufipes(in part)

104Femora and tibiae speckled …kingi

105Femora and tibiae with at most apical bands only …2

106Maxillary palpus shaggy; costa and vein 1 of wing without usual main dark areas …symesi

107Maxillary palpus smooth; 2nd main dark area of wing vein 1 well defined and with 2 pale interruptions …rufipes

108(in part)

109Maxillary palpus with 3 pale bands …2Maxillary palpus with 4 pale bands …6Maxillary palpus with apical 2 pale bands very broad, speckling on palpus segment 3; 2nd main dark area on wing vein 1 with 2 pale interruptions …stephensiMaxillary palpus with apical dark spot about equal to or longer than apical pale band; 2nd main dark area on wing vein 1 with 1 pale interruption.…33rd main dark area of wing vein 1 with a pale interruption, sometimes fused with preceding pale spot; scaling on abdomen very scanty, confined to tergum VIII or rarely VII …gambiae complex(in part)3rd main dark area of wing vein 1 without a pale interruption; abdominal terga fairly heavily clothed with cream or yellowish scales, especially on terga VI and VII …4Maxillary palpus speckled …hervyiMaxillary palpus not speckled ….5Foretarsomere 1 with some speckling; base of costa with 2 pale spots; stem of wing vein 2 entirely pale …salbaiiForetarsomere 1 not speckled; base of costa with 1 pale spot; stem of wing vein 2 extensively dark …dancalicusAll tarsi completely dark; wing without pale fringe spots posterior to vein 3 …vernus(in part)Tarsomeres 1 to 4 with conspicuous pale bands on at least the apices; wing with pale fringe spots up to lower branch of vein 5 or 6 …73rd main dark area of vein 1 with a pale interruption, sometimes fused with preceding pale area.…gambiae complex(in part)3rd main dark area without pale interruption …8Hindtarsomere 2 with about apical 0.4 to 0.5 white and the rest dark …multicinctusHindtarsomere 2 either with less than apical 0.4 white or else prominently marked with dark and pale bands …9Hindtarsomeres 2 to 4 with apical pale rings and otherwise dark except for 1 to 2 pale spots; no pale fringe spot opposite wing vein 6 …ardensisHindtarsomeres 2 to 4 with conspicuous dark and pale rings in addition to apical pale bands; pale fringe spot present opposite vein 6 …10Foretarsomeres mainly pale with narrow dark markings ….vinckeiForetarsomeres mainly dark with narrow pale rings …11Scales on abdominal tergum VIII dense and distributed over whole tergum, sometimes with a few scales on lateral borders of tergum VII …dureniScales on tergum VIII scanty and confined to posterior margin …millecampsi

110Maxillary palpus with 3 pale bands …2

111Maxillary palpus with 4 pale bands …6

112Maxillary palpus with apical 2 pale bands very broad, speckling on palpus segment 3; 2nd main dark area on wing vein 1 with 2 pale interruptions …stephensi

113Maxillary palpus with apical dark spot about equal to or longer than apical pale band; 2nd main dark area on wing vein 1 with 1 pale interruption.…3

1143rd main dark area of wing vein 1 with a pale interruption, sometimes fused with preceding pale spot; scaling on abdomen very scanty, confined to tergum VIII or rarely VII …gambiae complex

115(in part)

1163rd main dark area of wing vein 1 without a pale interruption; abdominal terga fairly heavily clothed with cream or yellowish scales, especially on terga VI and VII …4

117Maxillary palpus speckled …hervyi

118Maxillary palpus not speckled ….5

119Foretarsomere 1 with some speckling; base of costa with 2 pale spots; stem of wing vein 2 entirely pale …salbaii

120Foretarsomere 1 not speckled; base of costa with 1 pale spot; stem of wing vein 2 extensively dark …dancalicus

121All tarsi completely dark; wing without pale fringe spots posterior to vein 3 …vernus

122(in part)

123Tarsomeres 1 to 4 with conspicuous pale bands on at least the apices; wing with pale fringe spots up to lower branch of vein 5 or 6 …7

1243rd main dark area of vein 1 with a pale interruption, sometimes fused with preceding pale area.…gambiae complex

125(in part)

1263rd main dark area without pale interruption …8

127Hindtarsomere 2 with about apical 0.4 to 0.5 white and the rest dark …multicinctus

128Hindtarsomere 2 either with less than apical 0.4 white or else prominently marked with dark and pale bands …9

129Hindtarsomeres 2 to 4 with apical pale rings and otherwise dark except for 1 to 2 pale spots; no pale fringe spot opposite wing vein 6 …ardensis

130Hindtarsomeres 2 to 4 with conspicuous dark and pale rings in addition to apical pale bands; pale fringe spot present opposite vein 6 …10

131Foretarsomeres mainly pale with narrow dark markings ….vinckei

132Foretarsomeres mainly dark with narrow pale rings …11

133Scales on abdominal tergum VIII dense and distributed over whole tergum, sometimes with a few scales on lateral borders of tergum VII …dureni

134Scales on tergum VIII scanty and confined to posterior margin …millecampsi

135Wings entirely dark. or unicolorous …2Wings with at least some areas of paler scales on costa or vein 1, these being sometimes inconspicuous …5Maxillary palpus with 2 well-marked pale bands; hindfemur and hindtibia narrowly pale at apex …concolorMaxillary palpus and legs entirely dark …3Large species, wing length 4 mm or more …ruarinusSmall species, wing length 3.5 mm or less …4Very pale brown species with glossy scutum; semi-arid regions only ……rhodesiensis(in part)General coloration dark brown, scutum not so; cave-dwelling …caroni(in part)Maxillary palpus with 2 to 3 pale bands, pale at apex (sometimes indistinct) …6Maxillary palpus with or without pale bands, dark at apex ….9Erect head scales narrow, rod-like, all scales yellowish throughout; semi-arid regions only …dthaliErect head scales broader, scales white on vertex, dark laterally; all regions ….7Pale and dark areas on wing poorly contrasted; semi-arid regions only ….rhodesiensis(in part)Pale and dark areas on wing well contrasted …8Pale areas on wing very narrow, subcostal pale spot present on costa only; cave-dwelling …rodhainiPale areas on wing broader, subcostal pale spot on costa and vein 1 ……..rhodesiensis(in part)lounibosi Maxillary palpus with 3 pale bands, dark at apex …smithii(in part)Maxillary palpus unbanded or banding indistinct …10Cave-dwelling species; colour and contrast of dark and pale areas on wing variable …caroni(in part)hamoni vanhoofi Semi-arid regions; pale brown species with poorly contrasting light and dark areas on wing ….11 Erect head scales narrow, rod-like, all scales yellowish throughout ….azaniae(in part)Erect head scales broader, scales white on vertex, dark laterally...rhodesiensis(in part)

136Wings entirely dark. or unicolorous …2

137Wings with at least some areas of paler scales on costa or vein 1, these being sometimes inconspicuous …5

138Maxillary palpus with 2 well-marked pale bands; hindfemur and hindtibia narrowly pale at apex …concolor

139Maxillary palpus and legs entirely dark …3

140Large species, wing length 4 mm or more …ruarinus

141Small species, wing length 3.5 mm or less …4

142Very pale brown species with glossy scutum; semi-arid regions only ……rhodesiensis

143(in part)

144General coloration dark brown, scutum not so; cave-dwelling …caroni

145(in part)

146Maxillary palpus with 2 to 3 pale bands, pale at apex (sometimes indistinct) …6

147Maxillary palpus with or without pale bands, dark at apex ….9

148Erect head scales narrow, rod-like, all scales yellowish throughout; semi-arid regions only …dthali

149Erect head scales broader, scales white on vertex, dark laterally; all regions ….7

150Pale and dark areas on wing poorly contrasted; semi-arid regions only ….rhodesiensis

151(in part)

152Pale and dark areas on wing well contrasted …8

153Pale areas on wing very narrow, subcostal pale spot present on costa only; cave-dwelling …rodhaini

154Pale areas on wing broader, subcostal pale spot on costa and vein 1 ……..rhodesiensis

155(in part)

156lounibosi

157Maxillary palpus with 3 pale bands, dark at apex …smithii

158(in part)

159Maxillary palpus unbanded or banding indistinct …10

160Cave-dwelling species; colour and contrast of dark and pale areas on wing variable …caroni

161(in part)

162hamoni

163vanhoofi

164Semi-arid regions; pale brown species with poorly contrasting light and dark areas on wing ….11

165Erect head scales narrow, rod-like, all scales yellowish throughout ….azaniae

166(in part)

167Erect head scales broader, scales white on vertex, dark laterally...rhodesiensis

168(in part)

169Maxillary palpus shaggy to near tip …2Maxillary palpus smooth except at extreme base ….3Maxillary palpus entirely dark; hindtarsomeres 3 and 4 dark or narrowly pale at apices …obscurus(in part)Maxillary palpus with pale scales forming more or less definite pale bands; hindtarsomeres 3 and 4 narrowly or broadly pale at apices …tenebrosus(in part)Maxillary palpus with apex dark, sometimes only narrowly so …..4Maxillary palpus with apex pale …5Stem of wing vein 4 largely pale, upper branch of vein 5 with 2 pale spots or largely pale, fringe spots present opposite vein 4 and upper branch of vein 5 …tchekediiStem of wing vein 4 largely dark, upper branch of vein 5 with one narrow pale area, pale fringe spots absent...smithii(in part)Costa entirely dark except for a few indistinct pale scales subapically; maxillary palpus with a broad apical pale band and otherwise dark except for a narrow basal pale band …daudiOuter 0.5 of costa with 1–3 well-marked pale areas; maxillary palpus not so …6Maxillary palpus with 3 pale bands, subapical band broad and about equal in length to apical band ….7Maxillary palpus either with 4 pale bands or if with 3 bands then subapical band much shorter than the apical band …9Wing, apart from costa, generally very pale, basal 0.5 of stems of veins 2 and 4 entirely pale ….wellcomei(in part)Dark areas on wing greater than or about equal to pale areas, basal 0.5 of stems of veins 2 and 4 largely dark …8No pale fringe spots posterior to wing vein 3, stem of vein 5 pale except at fork and sometimes narrowly near base...erepensPale fringe spots present opposite all veins from wing apex to vein 5, stem of vein 5 broadly dark near base …keniensis(in part)Wing vein 5 entirely dark except for a single pale spot on the upper branch …fuscivenosus(in part)Wing vein 5 with extensive pale areas, upper branch of vein 5 with 2 pale spots ….10Hindtarsomeres 1 to 4 with distinct apical pale bands; scutum clothed with very narrow scales …distinctusHindtarsomeres 1 to 4 entirely dark or with a few pale scales at apices of 1 to 3; scutum scales broad …11Median scutal scales yellowish or bronze, white elsewhere …schwetzi(in part)Scutal scales white throughout...walravensi(in part)schwetzi (in part)

170Maxillary palpus shaggy to near tip …2

171Maxillary palpus smooth except at extreme base ….3

172Maxillary palpus entirely dark; hindtarsomeres 3 and 4 dark or narrowly pale at apices …obscurus

173(in part)

174Maxillary palpus with pale scales forming more or less definite pale bands; hindtarsomeres 3 and 4 narrowly or broadly pale at apices …tenebrosus

175(in part)

176Maxillary palpus with apex dark, sometimes only narrowly so …..4

177Maxillary palpus with apex pale …5

178Stem of wing vein 4 largely pale, upper branch of vein 5 with 2 pale spots or largely pale, fringe spots present opposite vein 4 and upper branch of vein 5 …tchekedii

179Stem of wing vein 4 largely dark, upper branch of vein 5 with one narrow pale area, pale fringe spots absent...smithii

180(in part)

181Costa entirely dark except for a few indistinct pale scales subapically; maxillary palpus with a broad apical pale band and otherwise dark except for a narrow basal pale band …daudi

182Outer 0.5 of costa with 1–3 well-marked pale areas; maxillary palpus not so …6

183Maxillary palpus with 3 pale bands, subapical band broad and about equal in length to apical band ….7

184Maxillary palpus either with 4 pale bands or if with 3 bands then subapical band much shorter than the apical band …9

185Wing, apart from costa, generally very pale, basal 0.5 of stems of veins 2 and 4 entirely pale ….wellcomei

186(in part)

187Dark areas on wing greater than or about equal to pale areas, basal 0.5 of stems of veins 2 and 4 largely dark …8

188No pale fringe spots posterior to wing vein 3, stem of vein 5 pale except at fork and sometimes narrowly near base...erepens

189Pale fringe spots present opposite all veins from wing apex to vein 5, stem of vein 5 broadly dark near base …keniensis

190(in part)

191Wing vein 5 entirely dark except for a single pale spot on the upper branch …fuscivenosus

192(in part)

193Wing vein 5 with extensive pale areas, upper branch of vein 5 with 2 pale spots ….10

194Hindtarsomeres 1 to 4 with distinct apical pale bands; scutum clothed with very narrow scales …distinctus

195Hindtarsomeres 1 to 4 entirely dark or with a few pale scales at apices of 1 to 3; scutum scales broad …11

196Median scutal scales yellowish or bronze, white elsewhere …schwetzi

197(in part)

198Scutal scales white throughout...walravensi

199(in part)

200schwetzi

201(in part)

202Maxillary palpus entirely dark or without distinct pale bands …2Maxillary palpus with 3 pale bands ….5Small, pale brown species, pale patches on wing indistinct, basal 0.25–0.5 of costa entirely dark; head scales narrow and yellowish ….azaniae(in part)Wing with well-contrasting pale and dark areas, basal 0.25 of costa with at least 1 pale area, even if narrow; head scales not so …3Costa with humeral pale spot, no subapical (preapical) pale spot on costa and vein 1 …obscurus(in part)Costa without a humeral pale spot, subapical pale spot present on costa and vein 1 …4Wing with pale fringe spots opposite all veins except vein 6 …jebudensisWing with no pale fringe spots posterior to vein 3 ….fainiWing generally pale, contrast between pale and dark areas, apart from costa and vein 1, poorly defined …turkhudiWing with well-contrasting pale and dark areas…62nd main dark area of wing vein 1 with 2 pale interruptions …72nd main dark area of wing vein 1 with at most 1 pale interruption …8Pale bands on maxillary palpus very narrow, at apices of segments 2 to 4 and not overlapping the joints; upper branch of wing vein 5 with a single pale spot …wilsoni(in part)Pale bands on maxillary palpus variable in width, distal 2 bands overlapping the joints; upper branch of wing vein 5 with 2 pale spots...rufipes(in part)Wing, apart from costa and vein 1, predominantly dark, no pale spots on basal 0.25 of costa …9Pale and dark areas on wing about equally distributed, humeral and presector pale spots present on costa …12Wing vein 6 dark …rageauiWing vein 6 with proximal pale spot …10Basal 0.2 of wing vein 1 either dark or with a proximal pale patch not extending to base.…smithii(in part)Basal 0.2 of wing vein 1 entirely pale ….11Wings scantily scaled, all wing scales very narrow.…fontinalisWings heavily scaled, upstanding scales moderately broad …lovettaeBasal pale band of maxillary palpus about equal to or slightly shorter than median band, broadly overlapping base of 3rd segment …cinereus(in part)Basal pale band of maxillary palpus either much shorter than median band, scarcely overlapping base of 3rd segment, or both basal and median pale bands very narrow …13Base of costa pale …multicolor(north-east Africa only)Base of costa dark...listeri(southern Africa only)azevedoi (south-western Africa only)seretsei (Botswana only)

203Maxillary palpus entirely dark or without distinct pale bands …2

204Maxillary palpus with 3 pale bands ….5

205Small, pale brown species, pale patches on wing indistinct, basal 0.25–0.5 of costa entirely dark; head scales narrow and yellowish ….azaniae

206(in part)

207Wing with well-contrasting pale and dark areas, basal 0.25 of costa with at least 1 pale area, even if narrow; head scales not so …3

208Costa with humeral pale spot, no subapical (preapical) pale spot on costa and vein 1 …obscurus

209(in part)

210Costa without a humeral pale spot, subapical pale spot present on costa and vein 1 …4

211Wing with pale fringe spots opposite all veins except vein 6 …jebudensis

212Wing with no pale fringe spots posterior to vein 3 ….faini

213Wing generally pale, contrast between pale and dark areas, apart from costa and vein 1, poorly defined …turkhudi

214Wing with well-contrasting pale and dark areas…6

2152nd main dark area of wing vein 1 with 2 pale interruptions …7

2162nd main dark area of wing vein 1 with at most 1 pale interruption …8

217Pale bands on maxillary palpus very narrow, at apices of segments 2 to 4 and not overlapping the joints; upper branch of wing vein 5 with a single pale spot …wilsoni

218(in part)

219Pale bands on maxillary palpus variable in width, distal 2 bands overlapping the joints; upper branch of wing vein 5 with 2 pale spots...rufipes

220(in part)

221Wing, apart from costa and vein 1, predominantly dark, no pale spots on basal 0.25 of costa …9

222Pale and dark areas on wing about equally distributed, humeral and presector pale spots present on costa …12

223Wing vein 6 dark …rageaui

224Wing vein 6 with proximal pale spot …10

225Basal 0.2 of wing vein 1 either dark or with a proximal pale patch not extending to base.…smithii

226(in part)

227Basal 0.2 of wing vein 1 entirely pale ….11

228Wings scantily scaled, all wing scales very narrow.…fontinalis

229Wings heavily scaled, upstanding scales moderately broad …lovettae

230Basal pale band of maxillary palpus about equal to or slightly shorter than median band, broadly overlapping base of 3rd segment …cinereus

231(in part)

232Basal pale band of maxillary palpus either much shorter than median band, scarcely overlapping base of 3rd segment, or both basal and median pale bands very narrow …13

233Base of costa pale …multicolor

234(north-east Africa only)

235Base of costa dark...listeri

236(southern Africa only)

237azevedoi

238(south-western Africa only)

239seretsei

240(Botswana only)

2413rd main dark area of wing vein 1 with a pale interruption.….23rd main dark area without a pale interruption..3Abdominal terga clothed with yellowish scales; hindtarsomeres 1 to 4 with broad apical pale bands.…christyiAbdominal terga without such scales; hindtarsomeres entirely dark or with a few pale scales at apices of hindtarsomeres 1 to 3 …schwetzi(in part)2nd main dark area of wing vein 1 with 2 pale interruptions…wilsoni(in part)2nd main dark area of wing vein 1 with 1 pale interruption …4Pale bands on maxillary palpus broad, basal band overlapping base of 3rd segment …cinereus(in part)Pale bands on maxillary palpus mostly narrow, basal band not overlapping base of 3rd segment ….5No pale fringe spots on wing posterior to vein 3; femora and tibiae inconspicuously speckled …..vernus(in part)Pale fringe spots on wing present opposite veins posterior to vein 3, sometimes including vein 6; femora and tibiae not speckled.…6Stem of wing vein 5 pale, at and adjacent to the fork …..garnhami(in part)Fork of wing vein 5 dark …7Wing length 4 mm or less; decumbent scutal scales not extending onto scutellum …demeilloni(Berg River form)Wing length 4.4 mm or more; some decumbent scales present on scutellum as well as scutum …carteri(in part)

2423rd main dark area of wing vein 1 with a pale interruption.….2

2433rd main dark area without a pale interruption..3

244Abdominal terga clothed with yellowish scales; hindtarsomeres 1 to 4 with broad apical pale bands.…christyi

245Abdominal terga without such scales; hindtarsomeres entirely dark or with a few pale scales at apices of hindtarsomeres 1 to 3 …schwetzi

246(in part)

2472nd main dark area of wing vein 1 with 2 pale interruptions…wilsoni

248(in part)

2492nd main dark area of wing vein 1 with 1 pale interruption …4

250Pale bands on maxillary palpus broad, basal band overlapping base of 3rd segment …cinereus

251(in part)

252Pale bands on maxillary palpus mostly narrow, basal band not overlapping base of 3rd segment ….5

253No pale fringe spots on wing posterior to vein 3; femora and tibiae inconspicuously speckled …..vernus

254(in part)

255Pale fringe spots on wing present opposite veins posterior to vein 3, sometimes including vein 6; femora and tibiae not speckled.…6

256Stem of wing vein 5 pale, at and adjacent to the fork …..garnhami

257(in part)

258Fork of wing vein 5 dark …7

259Wing length 4 mm or less; decumbent scutal scales not extending onto scutellum …demeilloni

260(Berg River form)

261Wing length 4.4 mm or more; some decumbent scales present on scutellum as well as scutum …carteri

262(in part)

2632nd and 3rd main dark areas of wing (median and preapical dark spots) absent from vein 1 …wellcomei(in part)2nd and 3rd main dark areas present on vein 1 …2Hindtarsomere 5 entirely pale, hindtarsomere 4 with broad apical and basal pale bands ….seydeliHindtarsomere 5 entirely dark, hindtarsomere 4 with narrow apical and basal pale bands.…3Upper branch of wing vein 5 with 1 pale spot, sometimes a vestigial 2nd pale spot …4Upper branch of wing vein 5 with 2 well-developed pale spots.…5Pale fringe spot present opposite wing vein 6; foretarsomeres 1 to 4 with conspicuous basal and apical pale bands …mortiauxiNo pale fringe spot opposite wing vein 6; foretarsomeres 1 to 4 narrowly pale apically only …bergheiSubapical pale band on maxillary palpus very narrow, confined to apex of 3rd segment …6Subapical pale band on maxillary palpus broad, overlapping apex of 3rd and base of 4th segment ….7Wing with base of costa with 2 pale interruptions ….brunnipesBasal 0.25 of costa entirely dark …walravensi(in part)Hindtarsomeres either all dark or with pale bands on tarsomeres 1 and 2 only ….8Hindtarsomeres 1 to 4 with well-marked apical pale bands …10Scutal fossae and lateral areas of scutum above wing root (supraalar area) without scales …..harperiScutal fossae and lateral areas of scutum above wing root (supraalar area) with scattered or abundant broadish scales ….9Subapical pale band on maxillary palpus about equal to or slightly shorter than apical band.…njombiensisSubapical pale band on maxillary palpus much narrower than apical band …walravensi(in part)Apical pale bands on hindtarsomeres 1 to 4 very broad, at least twice the apical width of the tarsomeres.…austeniiHindtarsomeres 1 to 4 with narrow pale bands, as long or shorter than the width of the tarsomeres …11Wing vein 3 largely dark or broadly dark at either end; scutal scales very narrow and golden …gibbinsi(in part)Wing vein 3 narrowly dark at ends; scutal scales various …12Scutal scales as in A ……hargreavesiScutal scales as in B …mousinhoiScutal scales as in C …marshalliiletabensis kosiensis hughi Scutal scales as in D …gibbinsi(in part)

2642nd and 3rd main dark areas of wing (median and preapical dark spots) absent from vein 1 …wellcomei

265(in part)

2662nd and 3rd main dark areas present on vein 1 …2

267Hindtarsomere 5 entirely pale, hindtarsomere 4 with broad apical and basal pale bands ….seydeli

268Hindtarsomere 5 entirely dark, hindtarsomere 4 with narrow apical and basal pale bands.…3

269Upper branch of wing vein 5 with 1 pale spot, sometimes a vestigial 2nd pale spot …4

270Upper branch of wing vein 5 with 2 well-developed pale spots.…5

271Pale fringe spot present opposite wing vein 6; foretarsomeres 1 to 4 with conspicuous basal and apical pale bands …mortiauxi

272No pale fringe spot opposite wing vein 6; foretarsomeres 1 to 4 narrowly pale apically only …berghei

273Subapical pale band on maxillary palpus very narrow, confined to apex of 3rd segment …6

274Subapical pale band on maxillary palpus broad, overlapping apex of 3rd and base of 4th segment ….7

275Wing with base of costa with 2 pale interruptions ….brunnipes

276Basal 0.25 of costa entirely dark …walravensi

277(in part)

278Hindtarsomeres either all dark or with pale bands on tarsomeres 1 and 2 only ….8

279Hindtarsomeres 1 to 4 with well-marked apical pale bands …10

280Scutal fossae and lateral areas of scutum above wing root (supraalar area) without scales …..harperi

281Scutal fossae and lateral areas of scutum above wing root (supraalar area) with scattered or abundant broadish scales ….9

282Subapical pale band on maxillary palpus about equal to or slightly shorter than apical band.…njombiensis

283Subapical pale band on maxillary palpus much narrower than apical band …walravensi

284(in part)

285Apical pale bands on hindtarsomeres 1 to 4 very broad, at least twice the apical width of the tarsomeres.…austenii

286Hindtarsomeres 1 to 4 with narrow pale bands, as long or shorter than the width of the tarsomeres …11

287Wing vein 3 largely dark or broadly dark at either end; scutal scales very narrow and golden …gibbinsi

288(in part)

289Wing vein 3 narrowly dark at ends; scutal scales various …12

290Scutal scales as in A ……hargreavesi

291Scutal scales as in B …mousinhoi

292Scutal scales as in C …marshallii

293letabensis

294kosiensis

295hughi

296Scutal scales as in D …gibbinsi

297(in part)

298Maxillary palpus with only apical pale band …gabonensisMaxillary palpus with 3 pale bands.….2Subapical pale band on maxillary palpus broad, about equal to or longer than apical dark band …3Apical dark band much longer than subapical pale band...17Apical pale band on hindtarsomere 4, and sometimes on hindtarsomeres 2 and 3, extending onto bases of succeeding tarsomeres ….4Bases of hindtarsomeres dark …72nd main dark area on wing vein 1 with 2 pale interruptions; bases of hindtarsomeres 4 and 5 broadly or narrowly pale …rufipes(in part)2nd main dark area on wing vein 1 with 1 pale interruption; bases of hindtarsomeres 4 and 5 at most narrowly pale …5Base of costa with 1 pale interruption, 3rd main dark area on costa and vein 1 much broader than subcostal pale spot …domicolusBase of costa with 2 pale interruptions, 3rd main dark area equal to or narrower than subcostal pale spot …6Pale fringe spot present opposite wing vein 6 …lloretiNo pale fringe spot opposite vein 6 …barberellusApices of hindtarsomeres 3 and 4 dark or at most with a few pale scales …8Apices of hindtarsomeres 1 to 3 and sometimes 4, distinctly pale banded …13Base of costa with 1 or no pale interruption …..9Base of costa with 2 pale interruptions…10Wing vein 6 either with pale fringe spot or with pale scales at apex of vein …brucei(in part)Wing vein 6 without pale fringe spot and no pale scales at apex …rivulorum(in part)Scutal scales fairly broad, extending over whole scutum and onto scutellum …carteri(in part)Scutal scales variable, but decumbent scales confined to at most anterior 0.66 of scutum …11Very small species, wing length 2.8 mm or less …brucei(in part)Small or moderate species, wing length 2.9 mm or more …12Hindtarsomeres entirely dark; preaccessory dark spot on wing vein 1 usually absent …freetownensisHindtarsomeres 1 and 2 narrowly but distinctly pale apically; preaccessory dark spot present on wing vein 1 …demeilloni(in part)3rd main dark area of costa equal to or shorter than subapical pale spot …flavicosta(in part)3rd main dark area much longer than subapical pale spot …14Scutal scales broadish and white, only slightly less dense on posterior 0.33 of scutum than anteriorly, and extending onto scutellum …flavicosta(in part)Scutal scales on posterior 0.33 of scutum scanty, narrow and yellowish-brown …15Moderate-sized species, wing length more than 3.2 mm …keniensis(in part)Small species, wing length 3.0 mm or less …16Foretarsomere 4 dark or indistinctly pale at apex; wing usually without pale fringe spot opposite vein 6 …..mouchetiForetarsomere 4 with well-marked apical pale band; wing with fringe spot opposite vein 6 …bervoetsi3rd main dark area of costa equal to or shorter than subapical pale spot …flavicosta(in part)3rd main dark area of costa much longer than subapical pale spot ….18Wing with fork of vein 5 pale..….garnhami(in part)Wing with fork of vein 5 dark …19Base of costa with 2 pale interruptions …demeilloni(in part)Base of costa with 1 or no pale interruption …20Small species, wing length about 2.4–3.3 mm …rivulorum(in part)Small or moderate-sized species, wing length 2.9–4.2 mm …demeilloni(in part)

299Maxillary palpus with only apical pale band …gabonensis

300Maxillary palpus with 3 pale bands.….2

301Subapical pale band on maxillary palpus broad, about equal to or longer than apical dark band …3

302Apical dark band much longer than subapical pale band...17

303Apical pale band on hindtarsomere 4, and sometimes on hindtarsomeres 2 and 3, extending onto bases of succeeding tarsomeres ….4

304Bases of hindtarsomeres dark …7

3052nd main dark area on wing vein 1 with 2 pale interruptions; bases of hindtarsomeres 4 and 5 broadly or narrowly pale …rufipes

306(in part)

3072nd main dark area on wing vein 1 with 1 pale interruption; bases of hindtarsomeres 4 and 5 at most narrowly pale …5

308Base of costa with 1 pale interruption, 3rd main dark area on costa and vein 1 much broader than subcostal pale spot …domicolus

309Base of costa with 2 pale interruptions, 3rd main dark area equal to or narrower than subcostal pale spot …6

310Pale fringe spot present opposite wing vein 6 …lloreti

311No pale fringe spot opposite vein 6 …barberellus

312Apices of hindtarsomeres 3 and 4 dark or at most with a few pale scales …8

313Apices of hindtarsomeres 1 to 3 and sometimes 4, distinctly pale banded …13

314Base of costa with 1 or no pale interruption …..9

315Base of costa with 2 pale interruptions…10

316Wing vein 6 either with pale fringe spot or with pale scales at apex of vein …brucei

317(in part)

318Wing vein 6 without pale fringe spot and no pale scales at apex …rivulorum

319(in part)

320Scutal scales fairly broad, extending over whole scutum and onto scutellum …carteri

321(in part)

322Scutal scales variable, but decumbent scales confined to at most anterior 0.66 of scutum …11

323Very small species, wing length 2.8 mm or less …brucei

324(in part)

325Small or moderate species, wing length 2.9 mm or more …12

326Hindtarsomeres entirely dark; preaccessory dark spot on wing vein 1 usually absent …freetownensis

327Hindtarsomeres 1 and 2 narrowly but distinctly pale apically; preaccessory dark spot present on wing vein 1 …demeilloni

328(in part)

3293rd main dark area of costa equal to or shorter than subapical pale spot …flavicosta

330(in part)

3313rd main dark area much longer than subapical pale spot …14

332Scutal scales broadish and white, only slightly less dense on posterior 0.33 of scutum than anteriorly, and extending onto scutellum …flavicosta

333(in part)

334Scutal scales on posterior 0.33 of scutum scanty, narrow and yellowish-brown …15

335Moderate-sized species, wing length more than 3.2 mm …keniensis

336(in part)

337Small species, wing length 3.0 mm or less …16

338Foretarsomere 4 dark or indistinctly pale at apex; wing usually without pale fringe spot opposite vein 6 …..moucheti

339Foretarsomere 4 with well-marked apical pale band; wing with fringe spot opposite vein 6 …bervoetsi

3403rd main dark area of costa equal to or shorter than subapical pale spot …flavicosta

341(in part)

3423rd main dark area of costa much longer than subapical pale spot ….18

343Wing with fork of vein 5 pale..….garnhami

344(in part)

345Wing with fork of vein 5 dark …19

346Base of costa with 2 pale interruptions …demeilloni

347(in part)

348Base of costa with 1 or no pale interruption …20

349Small species, wing length about 2.4–3.3 mm …rivulorum

350(in part)

351Small or moderate-sized species, wing length 2.9–4.2 mm …demeilloni

352(in part)

353Maxillary palpus with only apex pale…2Maxillary palpus with 3 pale bands.….5Base of costa with large (presector) pale spot, base of vein 1 pale …3Base of costa dark or with small pale spot, base of vein 1 dark …4Lower branch of wing vein 2 and upper branch of vein 4 with distinct pale spots …carnevaleiThese veins dark ….ovengensisSubapical pale spot on costa and wing vein 1 about as long as apical dark spot, fringe spots present opposite veins 3, lower branch of 4 and both branches of 5 …niliCongo formSubapical pale spot shorter, usually much shorter, than apical dark spot, no pale fringe spot opposite upper branch of vein 5 …nilisomalicus Hindtarsomeres 1 to 4 with pale bands overlapping the joints, at least hindtarsomere 5 pale basally …longipalpisPale banding on hindtarsomeres narrow and apical only …6Preaccessory dark spot on wing vein 1 about twice as long as pale spot on either side of it …fuscivenosus(in part)Preaccessory dark spot absent or, if present, shorter or only slightly longer than adjoining pale spots …7Basal area of wing vein 1 proximal to 1st main dark area, pale with a broad dark spot …culicifaciesBasal area of wing vein 1 entirely pale…8Subapical pale band on maxillary palpus longer than or equal to apical dark band AND 3rd main dark area of costa and vein 1 equal to or shorter than subapical pale spot ….aruniSubapical pale band on maxillary palpus much shorter than apical dark band, OR 3rd main dark area longer than subapical pale spot…9Moderate-sized species, wing length more than 3.3 mm …demeilloni(in part)Small species, wing length 3.2 mm or less …10Tip of wing vein 6 with a few pale scales, sometimes with fringe spot present …parensisTip of wing vein 6 dark with no fringe spot…funestus groupsergentii demeilloni (in part; mainly highlands)cameroni (extreme S. Africa only)

354Maxillary palpus with only apex pale…2

355Maxillary palpus with 3 pale bands.….5

356Base of costa with large (presector) pale spot, base of vein 1 pale …3

357Base of costa dark or with small pale spot, base of vein 1 dark …4

358Lower branch of wing vein 2 and upper branch of vein 4 with distinct pale spots …carnevalei

359These veins dark ….ovengensis

360Subapical pale spot on costa and wing vein 1 about as long as apical dark spot, fringe spots present opposite veins 3, lower branch of 4 and both branches of 5 …nili

361Congo form

362Subapical pale spot shorter, usually much shorter, than apical dark spot, no pale fringe spot opposite upper branch of vein 5 …nili

363somalicus

364Hindtarsomeres 1 to 4 with pale bands overlapping the joints, at least hindtarsomere 5 pale basally …longipalpis

365Pale banding on hindtarsomeres narrow and apical only …6

366Preaccessory dark spot on wing vein 1 about twice as long as pale spot on either side of it …fuscivenosus

367(in part)

368Preaccessory dark spot absent or, if present, shorter or only slightly longer than adjoining pale spots …7

369Basal area of wing vein 1 proximal to 1st main dark area, pale with a broad dark spot …culicifacies

370Basal area of wing vein 1 entirely pale…8

371Subapical pale band on maxillary palpus longer than or equal to apical dark band AND 3rd main dark area of costa and vein 1 equal to or shorter than subapical pale spot ….aruni

372Subapical pale band on maxillary palpus much shorter than apical dark band, OR 3rd main dark area longer than subapical pale spot…9

373Moderate-sized species, wing length more than 3.3 mm …demeilloni

374(in part)

375Small species, wing length 3.2 mm or less …10

376Tip of wing vein 6 with a few pale scales, sometimes with fringe spot present …parensis

377Tip of wing vein 6 dark with no fringe spot…funestus group

378sergentii

379demeilloni

380(in part; mainly highlands)

381cameroni

382(extreme S. Africa only)